1. Scottish Centre for Ecology and the Natural Environment, Institute of Biodiversity, Animal Health & Comparative Medicine, College of Medical, Veterinary and Life Sciences, University of Glasgow, Rowardennan G63 0AW, UK
2. Department of Animal and Plant Sciences, University of Sheffield, Sheffield S10 2TN, UK
3. Department of Biology, Section for Evolutionary Ecology, Lund University, SE-223 62 Lund, Sweden
4. Groningen Institute for Evolutionary Life Sciences (GELIFES), University of Groningen, 9747 AG Groningen, The Netherlands